• No results found

Future perspectives

How somatic and zygotic embryos develop in Pinus species is still poorly understood. However, the studies reported in this thesis are an important base for further research in embryology of Pinus. The transcriptome study provides an enormous source of data which can be used for further investigations of the molecular regulation of early embryo development. Especially interesting are the novel transcripts, i.e. transcripts with no homologue in Arabidopsis or other angiosperm species, which might be unique to species belonging to Pinus (or conifers). Functional studies of these genes would greatly expand our understanding of the regulation of embryo development in Pinus species.

Functional studies of the candidate genes, identified in paper (IV), are at present conducted in embryogenic cultures of Norway spruce. In the next step it will be important to perform functional studies in Scots pine. This requires that an efficient protocol for transforming embryogenic cultures of Scots pine is developed. None of the standard protocols used for transformation of conifers has so far been applicable to Scots pine.

Since there is a high risk of establishing cell lines giving rise to abnormal cotyledonary embryos when the embryogenic cultures are initiated from zygotic embryos at the stage of cleavage polyembryony, it would probably be better to initiate embryogenic cultures from more differentiated tissues. To accomplish this, further research is needed for elucidating how to regain embryogenic potential in older tissues.

References

Arya S, Kalia RK, Arya ID (2000) Induction of somatic embryogenesis in Pinus roxburghii Sarg.

Plant Cell Reports 19:775-780

Becwar MR, Nagmani R, Wann SR (1990) Initiation of embryogenic cultures and somatic embryo development in loblolly pine (Pinus taeda). Canadian Journal of Forest Research 20:810-817

Birol I, et al. (2013) Assembling the 20 Gb white spruce (Picea glauca) genome from whole- genome shotgun sequencing data. Bioinformatics pp 1-6

Bonga JM, Klimaszewska KK, von Aderkas P (2010) Recalcitrance in clonal propagation, in particular of conifers. Plant Cell Tissue and Organ Culture 100:241-254

Bozhkov PV, Ahn IS, Park YG (1997) Two alternative pathways of somatic embryo origin from polyembryonic mature stored seeds of Pinus koraiensis. Sieb et Zucc. Canadian Journal of Botany 75:509–512

Bozhkov PV, Suarez MF, Filonova LH, Daniel G, Zamyatnin AA, Rodriguez-Nieto S,

Zhivotovsky B, Smertenko A (2005) Cysteine protease mcII-Pa executes programmed cell death during plant embryogenesis. Proc Natl Acad Sci USA 102:14463-14468 Braybrook SA, Haranda JJ (2008) LECs go crazy in embryo development. Trends in Plant

Science 13:624-630

Buermans HPJ, den Dunnen JT (2014) Next generation sequencing technology: Advances and applications. Biochimica et Biophysica Acta 1842:1932-1941

Buchholz JT (1920) Embryo development and polyembryony in relation to the phylogeny of conifers. American Journal of Botany 7(4):125-145

Buchholz JT (1926) Origin of Cleavage Polyembryony in Conifers. Botanical Gazette 81(1):55- 71

Burg K, Helmersson A, Bozhkov P, von Arnold S (2007) Developmental and genetic variation in nuclear microsatellite stability during somatic embryogenesis in pine. Journal of Experimental Botany 58:687–698

Cairney J, Zheng L, Cowels A, Hsiao J, Zismann V, Liu J, Ouyang S, Thibaud-Nissen F, Hamilton J, Childs K, Pullman G.S, Zhang Y, Oh T, Buell CR (2006) Expressed Sequence Tags from loblolly pine embryos reveal similarities with angiosperm embryogenesis. Plant Molecular Biology 62:485-501

Carneros E, Celestino C, Klimaszewska K, Park Y-S, Toribio M, Bonga JM (2009) Plant regeneration in Stone pine (Pinus pinea L.) by somatic embryogenesis. Plant Cell Tissue and Organ Culture 98:165-178

Carney J, Pullman GS (2007) The cellular and molecular biology of conifer embryogenesis. New Phytologist 176:511-536

Cerda F, Aquea F, Gebauer M, Medina C, Arce-Johnson P (2002) Stable transformation of Pinus radiata embryogenic tissue by Agrobacterium tumefaciens. Plant Cell Tissue and Organ Culture 70:251-257

Chalupa V (1985) Somatic embryogenesis and plantlet regeneration from cultured immature and mature embryos of Picea abies /L./Karst. Communicationes Instituti Forestalis Czechosloveniae 14:57-63

Chen F, Dahal P, Bradford KJ (2001) Two Tomato Expansin Genes Show Divergent Expression and Localization in Embryos during Seed Development and Germination. Plant Physiology 127:928-936

Costa LM, Gutiérrez-Marcos JF, Dickinson HG (2004) More than a yolk: the short life and complex times of the plant endosperm. TRENDS in Plant Science 9(10):507-514 de Vega-Bartol JJ, Santos RR, Simões M, Miguel CM (2013) Normalizing gene expression by

quantitative PCR during somatic embryogenesis in two representative conifer species:

Pinus pinaster and Picea abies. Plant Cell Reports 32:715-729

Dogra PD (1967) Seed sterility and Disturbances in Embryogeny in Conifers with Particular Reference to Seed Testing and Tree Breeding in Pinaceae. Studia Forestalia Suecica Nr 45, Royal College of Forestry, Stockholm

Doyle J (1963) Proembryogeny in Pinus in Relation to That in Other Conifers: A Survey.

Proceedings of the Royal Irish Academy. Section B: Biological, Geological, and Chemical Science 62:181-216

Elbl P, Lira BS, Andrade SCS, Jo L, dos Santos ALW, Coutinho LL, Floh EIS, Rossi M (2014) Comparative transcriptome analysis of early somatic embryo formation and seed development in Brazilian pine, Araucaria angustifolia (Bertol.) Kuntze. Plant Cell Tissue and Organ Culture 120:903-915

Eriksson G, Ekberg I, Clapham D (2013). Genetics Applied to Forestry An Introduction. 3rd Edition. © Gösta Eriksson Inger Ekberg David Clapham, Elanders Sverige AB, Uppsala Sweden

Filonova LH, Bozhkov PV, Brukhin VB, Daniel G, Zhivotovsky B, von Arnold S (2000) Two waves of programmed cell death occur during formation and development of somatic embryos in the gymnosperm, Norway spruce. Journal of Cell Science 113(24):4399-4411

Filonova LH, von Arnold S, Daniel G, Bozhkov PV (2002) Programmed cell death eliminates all but one embryo in a polyembryonic plant seed. Cell Death & Differentiation 9:1057–

1062

Find J, Krogstrup P (2009) Integration of biotechnology, robot technology and visualization technology for development of methods for automated mass production of elite trees.

Working papers of the Finnish Forest Research Institute 114:72-77

Find JI, Hargreaves CL, Reeves CB (2014) Progress towards initiation of somatic embryogenesis from differentiated tissues of radiate pine (Pinus radiata D. Don) using cotyledonary embryos. In Vitro Cellular & Developmental Biology – Plant 50:190-198

Finer JJ, Kriebel HB, Becwar MR (1989) Initiation of embryogenic callus and suspension cultures of eastern white pine (Pinus strobus L.). Plant Cell Reports 8:203-206 Friedman WE, Carmichael JS (1998) Heterochrony and developmental innovation: evolution of

female gametophyte ontogeny in Gnetum, a highly apomorphic seed plant. Evolution 52(4):1016-1030

Garin E, Isabel N, Plourde A (1998) Screening of large numbers of seed families of Pinus strobus L. for somatic embryogenesis from immature and mature zygotic embryos. Plant Cell Reports 18:37-43

Gifford EM, Foster AS (1989) Morphology and Evolution of Vascular Plants. W.H. Freeman Company: New York. pp. 401-453

Grossnickle SC, Pait J (2008) Somatic embryogenesis tissue cultures for applying varietal forestry to conifer species. National Nursery Proceedings 2007. USDA Forest Service Proceedings RMRS-P-57, pp. 135-139

Grotkopp E, Rejánek M, Sanderson MJ, Rost TL (2004) Evolution of genome size in pines (Pinus) and its life-history correlates: super tree analyses. Evolution 58:1705-1729 Goldberg RB, de Paiva G, Yadegari R (1994) Plant embryogenesis: zygote to seed. Science

266:605-614

Hakman I, Fowke LC, von Arnold S, Erikson T (1985) The development of somatic embryos in tissue cultures initiated from immature embryos of Picea abies (Norway spruce). Plant Science 38:53-59

Hargreaves CL, Reeves CB, Find JI, Gough K, Josekutty P, Skudder DB, van der Maas SA, Sigley MR, Menzies MI, Low CB, Mullin TJ (2009) Improving initiation, genotype capture, and family representation in somatic embryogenesis of Pinus radiata by a combination of zygotic embryo maturity, media, and explant preparation. Canadian Journal of Forest Research 39:1566-1574

Hecht V, Vielle-Calzada JP, Hartog MV, Schmidt EDL, Boutilier K, Grossniklaus U, de Vries SC (2001) The Arabidopsis SOMATIC EMBRYOGENESIS RECEPTOR KINASE 1 Gene Is Expressed in Developing Ovules and Embryos and Enhances Embryogenic Competence in Culture. Plant Physiology 127:803-314

Helmersson A, von Arnold S, Kornel B, Bozhkov P (2004) High stability of nuclear

microsatellite loci during the early stages of somatic embryogenesis in Norway spruce.

Tree Physiology 24:1181-1186

Häggman H, Jokela A, Krajnakova J, Kauppi A, Niemi K, Aronen T (1999) Somatic

embryogenesis of Scots pine: cold treatment and characteristics of explants affecting induction. Journal of Experimental Botany 50(341):1769-1778

Högberg KA, Ekberg I, Norell L, von Arnold S (1998) Integration of somatic embryogenesis in a tree breeding progamme: a case study with Picea abies. Canadian Journal of Forest Research 28(10):1536-1545

Högberg KA, Bozhkov PV, von Arnold S (2003) Early selection improves clonal performance and reduces intraclonal variation of Norway spruce plants propagated by somatic embryogenesis. Tree Physiology 23:211-216

Ingouff M, Farbos I, Lagercrantz U, von Arnold S (2001) PaHB1 Is an Evolutionary Conserved HD-GL2 Homeobox Gene Expressed in the Protoderm During Norway Spruce Embryo Development. Genesis 30:220-230

Jacobs MR, Rubery PH (1988) Naturally Occuring Auxin Transport Regulators. Science 241(4863):346-9

Jain SM, Dong N, Newton RJ (1989) Somatic embryogenesis in Slash pine (Pinus elliotti) from immature embryos cultured in vitro. Plant Science 65:233-241

Jenik PD, Gillmor CS, Lukowitz W (2007) Embryonic Patterning in Arabidopsis thaliana. Annual Reviews Cell Developmental Biology 23:207-236

Jin F, Hu L, Yuan D, Xu J, Gao W, He L, Yang X, Zhang X (2012) Comparative transcriptome analysis between somatic embryos (SEs) and zygotic embryos in cotton: evidence for stress response functions in SE development. Plant Biotechnology Journal 12:161-173

Jones NB, van Staden J (1995) Plantlet Production from Somatic Embryos of Pinus patula. J Plant Physiology 145:519-525

Keinonen-Mettälä K, Jalonen P, Eurola P, von Arnold S, von Weissenberg K (1996) Somatic embryogenesis of Pinus sylvestris. Scandinavian Journal of Forest Research 11:242–

250

Klimaszewska K, Park Y-S, Overton C, Maceacheron I, Bonga JM (2001) Optimized somatic embryogenesis in Pinus strobus L. In Vitro Cell Developmental Biology – Plant 37:392-399

Klimaszewska K, Trontin JF, Becwar MR, Devillard C, Park YS, Lelu-Walter MA (2007) Recent progress in somatic embryogenesis of four Pinus spp. Tree and Forestry Science and Biotechnology 1:11-25

Klimaszewska K, Overton C, Stewart D, Rutledge RG (2010a) Initiation of somatic embryos and regeneration of plants from primordial shoots of 10 year-old somatic white spruce and expression profiles of 11 genes followed during the tissue culture process. Planta 233:635-647

Klimaszewska K, Pelletier G, Overton C, Stewart D, Rutledge RG (2010b) Hormonally regulated overexpression of Arabidopsis WUS and conifer LEC1 (CHAP3A) in transgenic white spruce: implications for somatic embryo development and somatic seedling growth.

Plant Cell Reports 29:723-734

Konar RN and Moitra A (1980) Ultrastructure, Cyto- and Histochemistry of Female Gametophyte of Gymnosperms. Gamete Research 3:67-97

Laine E, David A (1990) Somatic embryogenesis in immature embryos and protoplasts of Pinus caribaea. Plant Science 69:215-224

Lara-Chavez A, Flinn BS, Egertsdotter U (2011) Initiation of somatic embryogenesis from immature zygotic embryos of Oocarpa pine (Pinus oocarpa Schiede ex Schlectendal).

Tree Physiology 31:539-554

Larsson E, Sitbon F, Ljung K, von Arnold S (2008) Inhibited polar auxin transport results in aberrant embryo development in Norway spruce. New Phytologist 177:356–366 Le BH et al. (2010) Global analysis of gene activity during Arabidopsis seed development and

identification of seed-specific transcription factors. PNAS 107(18):8063-8070 Lelu-Walter MA, Bastien C, Drugeault A, Gouez ML, Klimaszewska K (1999) Somatic

embryogenesis and plantlet development in Pinus sylvestris and Pinus pinaster on medium with and without growth regulators. Physiologia Plantarum 105:719-728 Lelu-Walter MA, Bernier-Cardou M, Klimaszewska K (2006) Simplified and improved somatic

embryogenesis for clonal propagation of Pinus pinaster. Plant Cell Reports 25:767–

776

Lelu-Walter MA, Bernier-Cardou M, Klimaszewska K (2008) Clonal plant production from self- and cross-pollinated seed families of Pinus sylvestris (L.) through somatic

embryogenesis. Plant Cell Tissue and Organ Culture 92:31–45

Lelu-Walter MA, Thompson D, Harvengt L, Sanchez L, Toribio M, Pãques LE (2013) Somatic embryogenesis in forestry with a focus on Europe: state-of –the-art, benefits, challenges and future direction. Tree Genetics & Genomes 9:883-899

Lewis DR, Ramirez MV, Miller ND, Vallabhaneni P, Ray WK, Helm RF, Winkel BSJ, Muday GK (2001) Auxin and Ethylene Induce Flavonol Accumulation through Distinct Transcriptional Networks. Plant Physiology 156:144-164

Li XY, Huang FH (1996) Induction of somatic embryogenesis in loblolly pine (Pinus taeda L.).

In Vitro Cell Developmental Biology – Plant 32:129-135

Li XY, Huang FH, Gbur Jr EE (1998) Effect of basal medium, growth regulators and Phytagel concentration on initiation of embryogenic cultures from immature zygotic embryos of loblolly pine (Pinus taeda L.). Plant Cell Reports 17:298-301

Liao YK, Amerson HV (1995) Slash Pine (Pinus elliottii Engelm.) Somatic Embryogenesis I.

Initiation of Embryogenic Cultures from Immature Zygotic Embryos. New Forests 10:145-163

Lotan T, Ohto M, Yee KM, West MA, Lo R, Kwong RW, Yamagishi K, Fischer RL, Goldberg RB, Haranda JJ (1998) Arabidopsis LEAFY COTYLEDON1 is sufficient to induce embryo development in vegetative cells. Cell 93:1195-1205

Lu X, Chen D, Shu D, Zhang Z, Wang W, Klukas C, Chen L, Fan Y, Chen M, Zhang C (2013) The Differential Transcription Network between Embryo and Endosperm in the Early Developing Maize Seed. Plant Physiology 162:440-455

MacKay J, Becwar M, Park Y, Perfetti C, Cordero J, Pullman G, Lockhart L (2001) Genetics of somatic embryogenesis in loblolly pine. In: Dean JF (ed) Proceedings (publ no 48) 26th southern forest tree improvement Conference, University of Georgia, Athens, pp 40–47 MacKay JJ, Becwar MR, Park YS, Corderro JP, Pullman GS (2006) Genetic control of somatic

embryogenesis initiation in loblolly pine and implications for breeding. Tree Genetics

& Genomes 2:1-9

Maruyama E, Hosoi Y, Ishii K (2007) Somatic embryogenesis and plant regeneration in yakutanegoyou, Pinus armandii Franch. var. amamiana (Koidz.) Hatusima, an endemic and endangered species in Japan. In Vitro Cell Developmental Biology – Plant 43:28-34

Mathur G, von Arnold S, Nadgauda R (2000) Studies on somatic embryogenesis from immature zygotic embryos of chir pine (Pinus roxburghii Sarg.). Current Science 79(7):999-1004 Maximova SN, Florez S, Shen X, Niemenak N, Zhang Y, Curtis W, Guiltinan MJ (2014)

Genome-wide analysis reveals divergent patterns of gene expression during zygotic and somatic embryo maturation of Theobroma cacao L., the chocolate tree. BMC Plant Biology 14:185

Miguel C, Gonçalves, Tereso S, Marum L, Maroco J, Oliveira M.M (2004) Somatic

embryogenesis from 20 open-pollinated families of Portuguese plus trees of maritime pine. Plant Cell Tissue and Organ Culture 76:121-130

Mo LH, Egertsdotter U, von Arnold S (1996) Secretion of specific extracellular proteins by somatic embryos of Picea abies is dependent on embryo morphology. Annals of Botany 77:143-152

Montalbán IA, De Diego N, Moncaleán P (2012) Enhancing initiation and proliferation in radiate pine (Pinus radiata D. Don) somatic embryogenesis through seed family screening, zygotic embryo staging and media adjustments. Acta Physiologiae Plantarum 34:451-460

Mutz KO, Heilkenbrinker A, Lönne M, Walter JG, Stahl F (2013) Transcriptome analysis using next-generation sequencing. Current Opinion in Biotechnology 24:22-30

Nakazawa M, Yabe N, Ichikawa T, Yamamoto YY, Yoshizumi T, Hasunuma K, Matsui M (2001) DFL1, an auxin-responsive GH3 gene homologue, negatively regulates shoot cell elongation and lateral root formation, and positively regulates the light response of hypocotyl length. The Plant Journal 25(2):213-221

Nagmani R, Bonga JM (1985) Embryogenesis in sub-cultured callus of Larix decidua. Canadian Journal of Forest Research 15:1088-1091

Nagmani R, Diner AM, Sharma GC (1993) Somatic embryogenesis in longleaf pine (Pinus palustris). Canadian Journal of Forest Research 23:873-876

Niskanen AM, Lu J, Seitz S, Keinonen K, Von Weissenberg K, Pappinen A (2004) Effect of parent genotype on somatic embryogenesis in Scots pine (Pinus sylvestris). Tree Physiology 24(11):1259-65

North American Forest Commission (1998). North American Forest Commission, Nineteenth Session, Villahermosa, Mexico. [online] (1998) Available from

http://www.fao.org/UNFAO/Bodies/NAFC/nafc98/INF3-E.HTM [2016-01-04]

Nystedt B, et al. (2013) The Norway spruce genome sequence and conifer genome evolution.

Nature 497:579-584

Ohashi-Ito K, Bergmann DC (2006) Arabidopsis FAMA Controls the Final

Proliferation/Differentiation Switch during Stomatal Development. The Plant Cell 18:2493-2505

Owens JN, Morris SJ (1988) An Ultrastrucural Study of Fertilization in Douglas Fir [Pseudotsuga menziesii (Mirb.) Franco]. In: Cresti M et al. (eds.), Sexual Reproduction in Higher plants. © Springer-Verlag, Berlin Heidelberg

Park YS, Lelu-Walter MA, Harvengt L, Trontin JF, MacEacheron I, Klimaszewska K, Bonga JM (2006) Initiation of somatic embryogenesis in Pinus banksiana, P. strobus, P. pinaster, and P. sylvestris at three laboratories in Canada and France. Plant Cell Tissue and Organ Culture 86:87–101

Percy RE, Klimaszewska K, Cyr DR (2000) Evaluation of somatic embryogenesis for clonal propagation of western white pine. Canadian Journal of Forest Research 30:1867-1876

Pullman GS, Johnson S (2002) Somatic embryogenesis in loblolly pine (Pinus taeda L.):

improving culture initiation rates. Annals of Forest Science 59:663-668

Pullman G, Johnson S, Peter G, Cairney J, Xu N (2003) Improving loblolly pine somatic embryo maturation: comparison of somatic and zygotic embryo morphology, germination, and gene expression. Plant Cell Reports 21:747–758

Rao PS, Ozias-Akins P (1965) Plant Regeneration Through Somatic Embryogenesis in Protoplast Cultures of Sandalwood (Santalum album L.) Protoplasma 124:80-86

Raven PH, Evert RF, Eichhorn SE (1999) Biology of Plants, 6th Edition. © W.H. Freeman and Company/Worth Publishers. W.H. Freeman and Company, New York, U.S.A.

Reinert J (1959) Über die Kontrolle der Morphogenese und die Induktion von Adventivembryonen an Gewebekulturen aus Karotten. Planta 53:318-333

Roy Chowdhury C (1962) The embryology of conifers: a review. Phytomorphology 12:313-338 Sabala I, Elfstrand M, Farbos I, Clapham D, von Arnold S (2000) Tissue-specific expression of Pa18, a putative lipid transfer protein gene, during embryo development in Norway spruce (Picea abies). Plant Molecular Biology 42:461-478

Salajová T, Salaj J (1992) Somatic embryogenesis in European black pine (Pinus nigra Arn.).

Biologia Plantarum 34(3-4):213-218

Salajova T, Salaj J, Kormutak A (1999) Initiation of embryogenic tissues and plantlet regeneration from somatic embryos of Pinus nigra Arn. Plant Sciences 145:33-40 Salvo AAGD, Hirsch CN, Buell CR, Kaeppler SM, Kaeppler HF (2014) Whole Transcriptome

Profiling of Maize during Early Somatic Embryogenesis Reveals Altered Expression of Stress Factors and Embryogenesis-Related Genes. PLOS ONE 9:10

Schmeil O (1931) Leitfaden der Pflanzenkunde. Verlag von Quelle & LMeyer, Leipzig

Scheres B (2007) Stem-cell niches: nursey rhymes across kingdoms. Nature Reviews, Molecular Cell Biology. 8:345-354

Schoenbohm C, Martens S, Eder C, Forkmann G, Weisshaar B ( 2000) Identification of the Arabidopsis thaliana Flavonoid 3´-Hydroxylase Gene and Functional Expression of the Encoded P450 Enzyme. Biological Chemistry 381:749-753

Shendure J, Ji H (2008) Next-generation DNA sequencing. Nat Biotechnol 26(10):1135-45 Singh H (1978) Embryology of Gymnosperms. In: Zimmermann W, Carlquist Z, Ozenda P,

Wulff HD (eds) Handbuch der Pflanzenanatomie. Gebrüder Borntrager, Berlin, pp 187–241

Skinner D (1992) Ovule and embryo development, seed production and germination in orchard grown control pollinated loblolly pine (Pinus taeda L.) from coastal South Carolina.

Victoria, BC, Canada: Department of Biology, University of Victoria.

Skogssverige (2016). Svenska träd [online] (2016-01-04). Available from http://www.skogssverige.se /skog/svenska-trad [2016-01-04]

Smertenko AP, Bozhkov PV, Filonova LH, von Arnold S, Hussey PJ (2003) Reorganization of the cytoskeleton during developmental programmed cell death in Picea abies embryos.

Plant Journal 33:813-824

Smith SA, Beaulieu JM, Donoghue MJ (2010) An Uncorrelated Relaxed-Clock Analysis Suggests an Earlier Origin for Flowering Plants. Proceedings of the National Academy of Sciences 107(13):5897-5902

Steward FC, Mapes MO, Mears K (1958) Growth and organized development of cultured cells.

11. Organization from cultures grown from freely suspended cells. American Journal of Botany 45:705-708

Swedish Forest Agency (2014). Swedish Statistical Yearbook of Forestry. [online] (2014) Available from http://www.skogsstyrelsen.se/en/AUTHORITY/Statistics/Statistical-Yearbook-/Statistical-Yearbooks-of-Forestry/ [2016-01-04]

Tai HH, Tai GCC, Beardmore T (2005) Dynamic histone acetylation of late embryogenic genes during seed germination. Plant Molecular Biology 59:909-925

Tanaka M, Kikuchi A, Kamada H (2008) The Arabidopsis histone deacetylases HDA6 and HDA19 contribute to the repression of embryogenic properties after germination. Plant Physiology 146:149-161

ten Hove CA, Lu KJ, Weijers D (2015) Building a plant: cell fate specification in the early Arabidopsis embryo. Development 142:420-430

The Arabidopsis Genome Initiative (2000) Analysis of the genome sequence of the flowering plant Arabidopsis thaliana, Nature 408:796-815

To A (2006) A Network of Local and Redundant Gene Regulation Governs Arabidopsis Seed Maturation. THE PLANT CELL ONLINE 18(7):1642-1651

Tsuwamoto R, Yokoi S, Takahata Y (2010) Arabidopsis EMBRYOMAKER encoding an AP2 domain transcription factor plays a key role in developmental change from vegetative to embryonic phase. Plant Molecular Biology 73:481-492

Uddenberg D, Abrahamsson M, von Arnold S (2016) Overexpression of PaHAP3A stimulates differentiation of ectopic embryos from maturing somatic embryos of Norway spruce.

Tree Genetics & Genomes 12:18

Vestman D, Larsson E, Uddenberg D, Cairney J, Clapham D, Sundberg E, von Arnold S (2011) Important processes during differentiation and early development of somatic embryos of Norway spruce as revealed by changes in global gene expression. Tree Genetics &

Genomes 7:347-362

von Arnold S, Clapham D (2008) Spruce embryogenesis. In: Suárez MF, Bozhkov PV (eds) Plant embryogenesis: methods in molecular biology, Human Press, Totowa, NJ, 427:31–47 Wendrich JR, Weijers D (2013) The Arabidopsis embryo as a miniature morphogenesis model.

New Phytologist 199:14-25

Willemsen V, Bauch M, Bennet T, Campiho A, Wolkenfelt H, Xu J, Haseloff J, Scheres B (2008) The NAC Domain Transcription Factors FEZ and SOMBRERO Control the

Orientation of Cell Division Plane in Arabidopsis Root Stem Cells. Developmental Cell 15:913-922

Williams CG (2009) Conifer Reproductive Biology. Springer Science + Business Media B.V.

Wrickramasuriya AM, Dunwell JM (2015) Global scale transcriptome analysis of Arabidopsis embryogenesis in vitro. BMC Genomics 16:301

Yang M, Gao M, Yin X, Liu J, Xu Y, Zeng L, Li Q, Zhang S, Wang J, Zhang X, He Z (2013) Control of Rice Embryo Development, Shoot Apical Meristem Maintenance, and Grain Yield by Novel Cytochrome P450. Molecular Plant 6(6):1945-1960

Zhang Y, Zhang S, Han S, Li X, Qi L (2012) Transcriptome profiling and in silico analysis of somatic embryos in Japanese larch (Larix leptolepis). Plant Cell Reports 31:1637-1657 Zhu T, Moschou PN, Alvarez JM, Sohlberg JJ, von Arnold S (2014) WUSCHEL-RELATED

HOMEOBOX 8/9 is important for proper embryo patterning in the gymnosperm Norway spruce. Journal of Experimental Botany 65(22):6543-6552

Zhu T, Moschou PN, Alvarez JM, Sohlberg JJ, von Arnold S (2016) WUSCHEL-RELATED HOMEOBOX 2 is important for protoderm and suspensor development in the gymnosperm Norway spruce. BMC Plant Biology 16(1):19

Zimin A, et al. (2014) Sequencing and Assembly of the 22-Gb Loblolly Pine Genome. Genetics 196:875-890

Acknowledgements

I have met and worked with many fantastic people from all around the world during my years at the Plant Biology department. You have all together created such a nice working environment and directly or indirectly made it possible for me to write my thesis.

I would specially like to thank the following people:

First, I would like to thank my excellent and always optimistic supervisor Sara for giving me the opportunity to work together with her and with an interesting, however, challenging project. You are a fantastic woman and have been a true role model for me.

Secondly, I would like to thank my co-supervisors Eva, Jens, and Emma for taking the time to read and provide valuable comments on my manuscripts and thesis.

I would also like to thank all past and present members of the forest group:

Andreas, for all your help and support during the start of my PhD, for the movie nights, and for you just being a great fun guy. Daniel V, for being such a nice roommate and friend. It was great to start the PhD at the same time as you.

Emma, for becoming a supportive supervisor in the end of my PhD, but most of all for being such a nice and wonderful friend. You are always so smart and helpful! Daniel U, for letting me take part of your projects, for all the “endless”

interesting conversations and for baking delicious cookies!

Silvia and Irene, for struggling with the Pinus project together with me. I do not think I would have made it without the two of you. Silvia, I really miss you and I hope that we can meet again soon. Irene, it is great fun to work with you and you are always very helpful. Panos, for all the help with optimizing a

Related documents